Duvalius (Biharotrechus) - Naturhistorisches Museum Bern

Two new subterranean, microphthalmous
trechine beetles from the Mediterranean
area, and a synonymic note (Coleoptera:
Carabidae, Trechini)
Achille Casale
ABSTRACT
Contrib. Nat. Hist. 16: 1–16.
Duvalius (Biharotrechus) huberi sp. nov. from Southern Turkey (Toros chain, Geyik
Dağı Mt.), sampled at high altitude near the snow, is described and illustrated. Its
relationships are discussed: in particular, the new species seems to be the adelphotaxon of D. (B.) bortesii Casale & Vigna Taglianti, 1984, and forms with the latter
an isolated, Anatolian lineage within a Duvalius group widely spread in the Balkan
peninsula and the Carpathians.
Trechus (Trechus) messoulii sp. nov. from Northern Morocco (Rif massif, Taza,
Toghobeit cave), sampled in one of the deepest caves of Africa, is described and
illustrated. Its morphological features, both external and in male genitalia, are very
close to those of Trechus of the fulvus species group, and of Rifean species currently
attributed to genus Antoinella Jeannel.
The significance of these characters, in a both taxonomic and biogeographical context, is discussed, and the new synonymy is here proposed:
Antoinella Jeannel, 1937 = Trechus Clairville, 1806, new synonymy.
Keywords: Coleoptera, Carabidae, Trechini, Duvalius huberi, new species, Turkey, Trechus messoulii, new species, Morocco, Antoinella, new synonymy.
Introduction
The discovery of new subterranean taxa in the Euro-Mediterranean area is frequent, and their descriptions are a current, every-day taxonomical practice.
Nevertheless, some of these taxa are particularly interesting, because their
morphological features, and their geographical patterns of distribution, permit postulating the history and evolution of some lineages, and better understanding the taxonomic position of previously described relatives.
CONTRIBUTIONS TO NATURAL HISTORY
No. 16: 1–16 · 17 March 2011
This is the case of two new subterranean, microphthalmous trechine species here described, originating from Southern Turkey (Anatolia, Toros chain)
and Northern Morocco (Rif massif), respectively.
Material and methods
The following data come from two examined male individuals (see: Type
Material).
Male genitalia were dissected, dehydrated in ethanol, cleared in cold KOH,
and examined and illustrated, using standard techniques before being mounted on microscope slides attached to the respective specimens.
Line drawings were made using a camera lucida attached to stereomicroscopes Wild M-3 and Wild M-5, and a microscope Leitz Orthoplan.
Acronyms of measurements:
TL: body Total Length, from the anterior margin of clypeus to the apex of
elytra, measured along the suture.
L: overall Length, from apex of mandibles to apex of elytra, measured along
the suture.
PL/PW: ratio Length of Pronotum, as linear distance from the anterior to the
basal margin, measured along the midline/maximum Width of Pronotum,
as greatest transverse distance.
EL/EW: ratio Length of Elytra, as linear distance from the basal ridge to the
apex, measured along the suture/maximum Width of Elytra.
Collections:
NMBE Naturhistorisches Museum der Burgergemeinde Bern (Switzerland)
CCa
Casale collection, Torino (Italy)
Results
Duvalius (Biharotrechus) huberi sp. nov.
Type locality: Turkey, Southern Anatolia: Geyik Dağı.
Type material :
Holotype ♂ with the following data: "TR – Antalya Geyik Dağı 2600 mt, N 36°
54' E 32° 11' 22.5.2007 leg. C. Huber" (NMBE).
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Achille Casale
1 mm
Fig. 1: Duvalius (Biharotrechus) huberi sp. nov., male holotype, habitus, dorsal aspect.
New subterranean, microphthalmous trechine beetles from the Mediterranean area
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Diagnostic features
A medium sized (TL: 5.50 mm), glabrous, depigmented, microphthalmous
trechine species (Fig. 1), with the character states of the genus Duvalius Delarouzée, 1859 of Subgenus Biharotrechus Bokor, 1922 (Duvaliotes Jeannel,
1928, in the widest sense of Jeannel, 1928), Casale & Laneyrie (1982), and
Casale & Vigna Taglianti (1984).
Colour dark reddish. Pronotum cordiform, widened in front, with lateral sides constricted to the base. Elytra elongate, depressed, with humeral
angles rounded but evident. Chaetotaxy as in Fig. 1. Male genitalia as in Fig. 2.
Close to D. (Biharotrechus) bortesii Casale & Vigna Taglianti, 1984, from
which is distinguished by several different morphological features (see: Relationships).
Description
General features as in Fig. 1. Medium sized: TL: 5.50 mm ; L: 6.0 mm. Colour
dark reddish, with paler yellow reddish palpi, antennae and legs.
Dorsal surface moderately shiny; transversal cuticular microlines evident on
frons and pronotum, scarcely visible as polygonal meshes on elytral intervals.
Head elongate, with frontal furrows markedly impressed, angulate in the
anterior third, with evident transveral wrinkles; frons convex; supraorbital
setiferous punctures inserted on ranges slightly convergent backwards; eyes
small, as long as 1/4 of genae, flat, oblique and whitish, with slightly distinguishable ommatidia, pigmented only at margins; genae regularly curved,
convex, with deep transversal wrinkles, narrowed to the neck constriction;
labrum slightly emarginate at the anterior side; mentum tooth prominent, emarginate at apex; antennae elongate, exceeding backwards the level of the fourth
humeral setiferous puncture.
Pronotum cordiform transverse (PL/PW: 0.80), widest at the anterior third;
disc moderately convex; sides markedly arcuate in front, shortly and slightly
sinuate towards hind angles, which are evident but not prominent outside;
marginal furrows deep; anterior margin gently arcuate, front angles effaced;
base straight, slightly emarginate in the centre, fully beaded; basal foveae
deep.
Elytra elongate (EL/EW: 1.58), slightly widened at the apical third, much
wider than prothorax; disc depressed; humeri rounded but evident, with prehumeral margins oblique; lateral furrows relatively wide and deep; striae all
visible: striae 1–4 much deeper, 5–6 reduced to series of punctures, stria 7
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Achille Casale
Fig. 2: Duvalius (Biharotrechus) huberi sp. nov.,
male holotype, aedeagus
in left lateral aspect (a),
copulatory piece in dorsal aspect (b).
0.3 mm
b
a
almost effaced; apical striola very deep and curved, connected by some
points with apex of stria 5; intervals flat; apical carina developed. Chaetotaxy
as in Fig. 1: three setiferous discal punctures (two discal, one pre-apical) on
stria 3; umbilicate pores 5 and 6 very close to each other; apical and anguloapical punctures present, forming with the pre-apical puncture an apical triangle.
Legs long, slender; protibiae slightly dilated and sparsely pubescent at
apex, each with a long groove on the external side; protarsi in the male each
with two basal tarsomeres dilated and denticulate inwards.
Male genitalia as in Fig. 2. Median lobe of aedeagus relatively short (1.08
mm) and thickened, in lateral aspect regularly bent basally and slightly sinuate in the apical third, regularly narrowed at apex; basal bulb large, inflated,
with wide basal orifice; sagittal carina absent. Endophallus armed with an
elongate, "isotopic" copulatory piece, obliquely truncate at apex in lateral
aspect, regularly rounded in dorsal aspect, bifid at base, and a series of small,
sclerotized spines that fringe the basal tract of lateral sides of the copulatory
piece. Parameres robust, each with five apical setae.
Female genitalia: unknown.
New subterranean, microphthalmous trechine beetles from the Mediterranean area
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Specific epithet
I wish to dedicate this interesting new species to Dr. Charles Huber, curator of
the Entomological Department at the Berne Museum (Switzerland) and well
known specialist of carabids of the genus Nebria, who sampled with Dr. Werner
Marggi the holotype specimen, and offered it to me for study.
Geographical distribution and habitat
The only male individual of the new species here described was sampled on
the Northern slope of the Geyik Da ğı Mt. (Southern Anatolia, Toros chain,
Antalya) by Charles Huber and Werner Marggi at 2600 m of altitude, under
stones near the snow (Fig. 3). As many Duvalius species of the Euro-Mediterranean area, it is an upper hypogean species in the alpine zone.
Relationships
D. (Biharotrechus) huberi sp. nov. is particularly interesting for both taxonomical and biogeographical aspects.
First, from the taxonomical point of view, it is apparently the first adelphotaxon known so far of D. (Biharotrechus) bortesii Casale & Vigna Taglianti,
1984 – geographically vicariant in a massif (Bey Dağları Mt., Muğla province)
at 80 kms SW of Geyik Dağı – to which it seems related by many morphological features, and the peculiar shape of both median lobe of aedeagus and
copulatory piece. From the latter, D. (Biharotrechus) huberi sp. nov. is distinguished by the more elongate head, the wider pronotum with basal angles
smaller and not prominent; the shorter, more depressed elytra, with shallower
and less deeply punctuate striae; and the different shape of aedeagus: in D.
(B.) huberi sp. nov. the median lobe is shorter and more thickened, and endophallus has reduced sclerotized spines and scales, and much more elongate
copulatory piece. It should also be recalled that individuals of D. (B.) bortesii
were sampled in a pit at 1200 m, whereas D. huberi sp. nov. in the alpine zone
at 2600 m near the snow. But this datum is scarcely informative: in fact, species with this degree of low specialization to the subterranean environment
can be found both in caves at lower altitude and in the shallow hypogean zone
at high altitude.
Validity and limits of the Subgenus Duvaliotes Jeannel, 1928, and its distinction from the Subgenus Duvalidius Jeannel, 1928, currently treated as
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Achille Casale
a
b
Fig. 3: Duvalius (Biharotrechus) huberi sp. nov., type locality: Geyik Daği Mt. (Southern Anatolia,
Toros chain, Antalya) (a); detail of the habitat (b) (photo C. Huber).
New subterranean, microphthalmous trechine beetles from the Mediterranean area
7
synonyms of Biharotrechus Bokor, 1922, and Hungarotrechus Bokor, 1922,
respectively (see Moravec & al. 2003, and Lorenz 2005) have been widely
debated by Casale & Vigna Taglianti (1984). What is evident, is the fact that
D. (B.) huberi sp. nov. and D. (B.) bortesii (the latter, described as Duvaliotes)
form a homogeneous species group, and are so far the only representatives
in Southern Anatolia of a lineage widely spread in the Carpathians, the Balkan
peninsula and the Dinaric Alps.
Therefore, as shown by Casale & Vigna Taglianti (1999), in Anatolia the
genus Duvalius includes so far representatives of some different lineages:
in the central and eastern Pontic chain, and in eastern Anatolia, some forest
dweller or troglophilic species, all probably belonging to the Caucasian antoniae species group. Markedly isolated seem to be D. bicikensis Perrault, 1971,
from the Giresun Dağları, apparently related to the Eastern-European Hungarotrechus (Duvalidius of authors) complex, and D. bruschii Vigna Taglianti, 1999,
probably related to the southern Balkan krueperi-group. By contrast, in the
Western Toros, the genus is represented by three peculiar, subterranean but
oculate species, with very different relationships: D. bortesii and D. huberi sp.
nov., treated in the present contribution, are apparently related to the Balkan
Biharotrechus lineage; and D. huetheri Jeannel, 1934, present in several caves
of Antalya and Muğla provinces, redescribed with different names, is related
to the "Southern Aegean" lineage of D. diaphanus Rottenberg, 1874, and D.
mirei Deuve, 2001, both endemic to Mt. Taygetos (Greece, Peloponnesus).
Trechus (Trechus) messoulii sp. nov.
Type locality: Morocco, Rif (Taza): Toghobeit cave. Type material:
Holotype ♂ with the following data: "Maroc, Rif, Gr. Toghobeit, 25–30.VII.
1999 leg. M. Messouli" (CCa).
Diagnostic features
A small-medium sized (TL: 4.25 mm), glabrous, depigmented, microphthalmous trechine species (Fig. 4), with the character states of the genus Trechus
Clairville, 1806 of the fulvus species group in the sense of Jeannel (1927),
Casale (1983), Ortuño (2008), Reboleira & al. (2010), and in the widest sense
that will be discussed in Conclusions.
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Achille Casale
1 mm
Fig. 4: Trechus (Trechus) messoulii sp. nov., male holotype, habitus, dorsal aspect.
New subterranean, microphthalmous trechine beetles from the Mediterranean area
9
Colour dark yellow reddish. Pronotum markedly transverse, widened
in front, with lateral sides regularly curved, slightly constricted and sinuate
towards hind angles, which are obtuse. Elytra elongate-ovate, widened in the
apical third, depressed, with humeral angles rounded. Chaetotaxy as in Fig. 4.
Male genitalia as in Fig. 5.
Close to some North African species attributed to the genus Antoinella
Jeannel, 1937, in the sense of Casale (1982), Mateu & Comas (2006) and
Mateu & Escolà (2006), from which it is recognized by several different morphological features (see: Relationships).
Description
General features as in Fig. 4. Small-medium sized: TL: 4.25 mm; L: 4.50 mm.
Colour uniformly dark yellow reddish. Dorsal surface moderately shiny; transversal cuticular microlines evident on frons and pronotum, scarcely visible as
polygonal meshes on elytral intervals.
Head robust, stout, with frontal furrows markedly impressed, angulate in
the anterior third; frons convex; supraorbital setiferous punctures inserted
on ranges slightly convergent backwards; eyes very small, as long as 1/5 of
genae, flat, oblique and whitish, with slightly distinguishable ommatidia, pigmented only at margins; genae regularly curved, slightly convex; neck wide,
neck constriction moderate; labrum slightly emarginate at the anterior side;
mentum tooth slightly scooped-out distally; antennae relatively long, exceeding backwards the level of the fourth humeral setiferous puncture.
Pronotum markedly transverse (PL/PW: 0.74), widened in front; disc convex; sides regularly curved, slightly constricted and shortly sinuate towards
hind angles, which are obtuse; marginal furrows narrow and shallow; anterior
margin almost straight, front angles effaced; base slightly oblique at sides,
not beaded; basal foveae deep; basal area densely wrinkled.
Elytra elongate-ovate (EL/EW 1.47), widened in the apical third, depressed,
with humeral angles rounded, widened at the apical third, much wider than
prothorax; disc depressed; lateral furrows wide and deep; striae all complete
and visible, moderately impressed, punctate; apical striola curved and connected with apex of stria 5; intervals flat; apical carina slightly prominent.
Chaetotaxy as in Fig. 4: three setiferous discal punctures (two discal on stria 3,
one pre-apical on apex of interval 3); umbilicate pores 5 and 6 very close to
each other; apical and angulo-apical punctures present, forming with the preapical puncture an apical triangle.
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Achille Casale
Fig. 5: Trechus (Trechus)
messoulii sp. nov., male
holotype, aedeagus in
left lateral aspect (a)
and in dorsal aspect (b).
0.3 mm
a
b
Legs long, slender; femora robust, thickened; protibiae dilated, very shallowly grooved on the external side, and sparsely pubescent at apex; protarsi
in the male each with two basal tarsomeres dilated and denticulate inwards.
Male genitalia as in Fig. 5. Median lobe of aedeagus elongate (1.05 mm
long, measured from apex to the posterior margin of basal bulb, basal carina
excluded), slender, slightly thickened in the middle, markedly narrowed and
curved on the dorsal side at apex, with apical lamina asymmetrical widened
and ventrally hooked in lateral aspect; basal bulb with big, rounded sagittal
carina, not extended on the ventral side of the bulb. Endophallus with complex structure: in left lateral view, it shows a dorsal group of scales that covers a sclerotized piece of subtriangular form; in the inner part, it shows a second sclerotized piece, covered with scarcely sclerotized scales, elongate, narrowed and bent distally. Parameres robust, asymmetrical; left paramere with
an elongate, ventral basal lobe on the ventral side (as is typical in Antoinella
species: see below), with five apical setae; right paramere shorter, with three
apical setae.
Female genitalia: unknown.
New subterranean, microphthalmous trechine beetles from the Mediterranean area
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Specific epithet
I wish to dedicate this interesting new species to Dr. Mohamed Messouli,
active speleologist in Marrakesh, who sampled the holotype specimen several
years ago, during an expedition with the Spéléo Club of Blois (France), and
through our friend Lucien C. Genest offered it to me for study.
Geographical distribution and habitat
The male holotype of this new species was sampled in the cave Toghobeit
(Northern Morocco, Rif Massif, Taza, near Chaouen or Chefchaouen). This subterranean system is so far one of the deepest caves of the African continent:
located at 1700 m of altitude, its actual depth is -732 m, and the development
reaches 3050 m (after: "Inventaire Spéléologique du Maroc", 1981: DC 5, Kef
Toghobeit). The only examined individual was collected in the "salle Renner"
at -120 m depth, but a second individual (not available at the moment) was
collected in the "salle du bivouac" and entrusted to L. C. Genest (Messouli,
personal communication).
Relationships
Trechus messoulii sp. nov., for both its external features and peculiar characters of male genitalia – in particular, the structure of copulatory pieces in
endophallus (a sclerotized piece of triangular shape, a second elongate piece,
and some groups of sclerotized scales) – is close to some Trechus species of
the fulvus species group in the sense of Reboleira & al. (2010) (in particular,
for the widened and hooked apical lamina of the median lobe of the aedeagus,
it recalls T. arribasi Jeanne, 1988, endemic to the Northern Iberian peninsula).
But it is mostly related to Algerian species (Jeannel, 1927; Casale, 1983;
Ortuño, 2008) and several subterranean North African species of the Rif massif
(Morocco) currently treated as Antoinella Jeannel, 1937 (Casale, 1982; Mateu
& Comas, 2006; Mateu & Escolà, 2006). In particular, it seems to be close to A.
iblanensis Mateu & Escolà, 2006, and A. fadriquei Mateu & Escolà, 2006. It
is, however, markedly distinct by the smaller size (5.8–5.9 mm in A. iblanensis
and A. fadriquei, respectively, about 4.5 mm in T. messoulii sp. nov.), and the
shape of the median lobe of the aedeagus, which in T. messoulii sp. nov. is
slender and elongate as in A. iblanensis, but shows an apical lamina similar to
that of A. fadriquei, more curved dorsally and more obtusely truncate.
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Achille Casale
In this case, the combination of both morphological features and biogeographical considerations allows the following remarks:
1. From the taxonomical point of view, the so-called genus Antoinella Jeannel, 1937, appears as a paraphyletic unit, without any relationships with
the genus Duvalius. This datum is evident from both morphological features, as already supposed by Jeannel (1937) in the original description
of the genus, and supported by Casale (1982, 1983); furthermore, it has
been confirmed by a recent molecular phylogeny (Faille & al. 2010). Therefore, either it will be demonstrated that Trechus is a polyphyletic unit, and in
this case the T. fulvus species group should merit the genus rank (named
Antoinella), or the latter is a simple lineage within this group, and an
objective junior synonym of Trechus (in the widest sense).
2. The only current distinctive character between Antoinella and Trechus
of the fulvus species group, i.e. protibiae pubescent in Antoinella (as
in Duvalius species), and glabrous in Trechus, is not a constant diagnostic feature: Reboleira & al. (2010) for instance, who have just described
T. tatai from Portugal, specified that it has "conspicuously hairy (including the protibiae)" legs. Furthermore, the structure of the median lobe of
aedeagus, endophallus, and asymmetrical parameres (the left one with
an elongate, ventral basal lobe on the ventral side) is absolutely the same
both in Antoinella species and in some Algerian Trechus of the fulvus species group, as illustrated by Jeannel (1927) and Casale (1983).
3. Finally Ortuño (2008) and Reboleira & al. (2010), in their excellent contributions, exhaustively showed that the T. fulvus species group, in its wide
distribution along North Africa, the Iberian Peninsula and, thanks to the
wide distribution of T. fulvus, also along the Atlantic coast of northern Europe (Jeannel 1927), is heterogeneous, and includes at least two lineages.
In this context, it is evident that the so-called Antoinella species, with the
martinezi-lineage (sensu Ortuño & Arillo 2005; Ortuño 2008), form a typical
subterranean lineage within the Trechus fulvus species group, which has survived in the south-western Mediterranean region. The present distribution of
extant species suggests a Betic-Rifean origin (see also Ortuño 2008: fig. 4),
with strong biogeographical coherence: a well known case in entomological
literature. Another interesting fact is the surprising speciosity of this group in
the Rif massif: in fact every cave, so far, revealed a separate species markedly distinct from its relatives, all with markedly reduced eyes, depigmented
integument, and other troglomorphic features.
New subterranean, microphthalmous trechine beetles from the Mediterranean area
13
In conclusion, the following new synonymy is proposed:
Antoinella Jeannel, 1937 (type species: Duvalius groubei Antoine, 1935)
junior synonym of Trechus Clairville, 1806 (type species : Carabus rubens
Fabricius sensu Clairville, 1806 = Carabus quadristriatus Schrank, 1781)
A separate question is the validity and rank of Irinea Mateu & Comas,
2006, described as subgenus of Antoinella. The type species, A. (I.) aurouxi
Mateu & Comas, 2006, shows marked distinctive features: in particular, only
one basal tarsomere is dilated in fore tarsi of males, median lobe of aedeagus
without basal sagittal carina – typically present in all "Antoinella" species
and Trechus of the fulvus species group – and endophallus of simple structure,
with only one small copulatory piece.
Nevertheless, these characters are not unique and peculiar to Irinea, but
present in several species attributed to Trechus in the current (and widest)
sense. In a contribution dedicated to the Ethiopian Trechus species, Ortuño &
Novoa (in press) discuss the sexual dimorphism in species of this genus, and
stress that the occurrence of a single dilated articulation on the protarsus in
males is present in the Nearctic subgenus Microtrechus Jeannel, 1927, in Trechus (Atlantotrechus) cautus Wollaston, 1854, from Madeira and in some
Ethiopian species (Ortuño, personal communication; Pawlowski 2003). Furthermore, it must be recalled that recently Vigna Taglianti & Magrini (2009)
described the subgenus Minitrechus for Trechus gypaeti, new species from
Ethiopia, in which only one basal tarsomere is dilated in fore tarsi of males.
Therefore, given that the apomorphy of a single dilated tarsomere in males
appears sporadically in Trechus belonging to different lineages and in different areas, this character seems not a valid characteristic able to distinguish
just one group within Trechus in the widest sense (Ortuño & Novoa, in press;
Ortuño, personal communication).
Therefore taxonomic position, validity, and phylogenetic relationships of
Irinea merit further investigation.
Acknowledgements
For the material and information on which the present descriptions are based,
I am particularly indebted to my colleagues Charles Huber (Bern, Switzerland)
and Werner Marggi (Thun, Switzerland), and Mohamed Messouli (Marrakesh,
Morocco). For notes, suggestions and information on Trechus of the fulvus species group, I am very grateful to Vicente Ortuño (Madrid, Spain), well known
specialist of Trechini of the Iberian Peninsula and Northern Africa, who kindly
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Achille Casale
and amicably authorized me to anticipate some results of a not yet published
contribution. Finally, for their constant help in the laboratory, I wish to thank
my collaborators Ivo Manca and Paolo Marcia (Sassari, Italy).
References
Casale, A. (1982): Nuovi Carabidi di Marocco, di Grecia e di Papua - Nuova Guinea (������
Coleoptera) — Revue suisse de Zoologie 89: 229–244.
Casale, A. (1983): Un nuovo Trechus d’Algeria (Coleoptera, Carabidae) — Bollettino del
Museo regionale di Scienze naturali, Torino 1(1): 121–126.
Casale, A. & Laneyrie, R. (1982): Trechodinae et Trechinae du monde. Tableau des sousfamilles, tribus, séries phylétiques, genres, et catalogue général des espèces — Mémoires
de Biospéologie 9: 226 pp.
Casale, A. & Vigna Taglianti, A. (1984): Due nuovi Carabidi cavernicoli dell’Anatolia meridionale (Coleoptera, Carabidae) — Fragmenta entomologica (Roma) 17: 309–329.
Casale, A. & Vigna Taglianti, A. (1999): Caraboid beetles (excl. Cicindelidae) of Anatolia,
and their biogeographical significance (Coleoptera, Caraboidea) — Biogeographia 20:
277–406.
Faille, A., Casale, A. & Ribera, I. (2010): Phylogenetic relationships of Western Mediterranean subterranean Trechini groundbeetles (Coleoptera: Carabidae) — Zoologica Scripta
(printed, version on line).
Jeannel, R. (1927): Monographie des Trechinae 2. Morphologie comparée et distribution
géographique d'un group de Coléoptères — L’Abeille, Paris 33: 1–592.
Jeannel, R. (1928): Monographie des Trechinae 3. Morphologie comparée et distribution
géographique d’un groupe de Coléoptères — L’Abeille, Paris 35: 1–808.
Jeannel, R. (1937): Nouveaux Trechinae paléarctiques — Bulletin de la Société entomologique de France 42: 82–88.
Lorenz, W. (2005): Systematic list of extant ground beetles of the world (Insecta Coleoptera "Geadephaga": Trachypachidae and Carabidae. incl. Paussinae, Cicindelinae, Rhysodinae). Tutzing, iv + 530 pp.
Mateu, J. & Comas, J. (2006): Una nueva especie del género Antoinella Jeannel, 1937 de
Marruecos (Coleoptera: Carabidae: Trechinae) — Heteropterus Revista de Entomologia 6:
15–18.
Mateu, J. & Escolà, O. (2006): El género Antoinella Jeannel, 1937 (Coleoptera, Carabidae,
Trechinae) tres especies nuevas de Marruecos: A. espanyoli sp. n., A. iblanensis sp. n. y
A. fadriquei sp. n. — Animal Biodiversity and Conservation 29(2): 117–121.
Moravec, P., Uéno, S.-I. & Belousov, I.A. (2003): Carabidae: Trechinae: Trechini. — In: Löbl,
I. & Smetana, A. (Eds.), Catalogue of Palaearctic Coleoptera, Vol. 1., pp. 288–346, Apollo
Books, Stenstrup.
Ortuño, V.M. (2008): Taxonomy and Systematics of a Hypogean Trechine from Southern
Spain: Trechus breuili Jeannel (Coleoptera: Carabidae) — The Coleopterists Bulletin 62 (4):
501–507.
Ortuño, V.M. & Arillo, A. (2005): Description of a new hypogean species of the genus Trechus Clairville, 1806 from eastern Spain and comments on the Trechus martinezi-lineage
(Coleoptera: Adephaga: Carabidae) — Journal of Natural History, 39(40), 3483–3500.
New subterranean, microphthalmous trechine beetles from the Mediterranean area
15
Ortuño, V.M. & Novoa, F. (in press): A new species of Trechus from the Ethiopian Highlands
(Coleoptera: Carabidae: Trechinae) and Key to the Trechus Species of Ethiopia — Annals of
the Entomological Society of America.
Pawlowsky, J. (2003): Preliminary report on Ethiopian species of the genus Trechus Clairville, 1806 (Coleoptera: Carabidae) — Baltic Journal of Coleopterology 3(2): 157–160.
Reboleira, A.S., Ortuño, V.M., Gonçalvesi, F. & Oromi, P. (2010): A hypogean new species
of Trechus Clairville, 1806 (Coleoptera, Carabidae) from Portugal and considerations about
the T. fulvus species group — Zootaxa 2689: 15–26.
Vigna Taglianti, A. & Magrini, P. (2009): A new subgenus and a new species of Trechus from
Ethiopia (Coleoptera, Carabidae) — Fragmenta entomologica (Roma) 41(1): 69–76.
Address of the author:
Prof. Dr. Achille Casale
Università di Sassari
Dipartimento di Zoologia e Genetica Evoluzionistica
Via Muroni 25
I – 07100 Sassari (Italy)
E-mail: casale@uniss.it; a_casale@libero.it
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Achille Casale